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Type: Articles
Published: 2007-12-11
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A revision of Entobdella Blainville in Lamarck, 1818, with special emphasis on the nominal (type) species “Entobdella hippoglossi (Müller, 1776) Blainville, 1818” (Monogenea: Capsalidae: Entobdellinae) from teleost flatfishes, with descriptions of three new species and a new genus

School of Biological Sciences, University of East Anglia, Norwich, NR4 7TJ, Norfolk, U.K.
Monogenean Research Laboratory, Parasitology Section, The South Australian Museum, North Terrace, Adelaide, South Australia 5000, Australia Marine Parasitology Laboratory, School of Earth and Environmental Sciences (DX 650 418), The University of Adelaide, North Terrace, Adelaide, South Australia 5005, Australia
School of Biological Sciences, University of East Anglia, Norwich, NR4 7TJ, Norfolk, U.K.
Platyhelminthes Monogenea Capsalidae Entobdellinae Entobdella brinkmanni n. sp. Entobdella hippoglossi sensu stricto Entobdella stenolepis n. sp. Entobdella vanbenedeni n. sp. Branchobdella n. gen. Branchobdella pugetensis n. comb. teleost flatf


Species of Entobdella Blainville in Lamarck, 1818, from skin (less commonly gills) of teleost flatfishes and occasionally from round-bodied teleosts are reviewed, with special reference to the nominal (type) species “Entobdella hippoglossi (Müller, 1776) Blainville, 1818” (Monogenea: Capsalidae: Entobdellinae) and to E. pugetensis Robinson, 1961. In the late 20th century the following 5 species were synonymised with E. hippoglossi: E. brattstroemi Brinkmann, 1952; E. curvunca Ronald, 1957; E. rosaceus Crane, 1972; E. squamula (Heath, 1902) Johnston, 1929; E. steingroeveri (Cohn, 1916) Johnston, 1929. We have shown that E. brattstroemi and E. squamula are valid, E. curvunca is a synonym of E. hippoglossi and E. rosaceus and E. steingroeveri are species inquirendae. Specimens of E. steingroeveri, first described in 1916 by Cohn, have been re-examined. Parasites from the Pacific Ocean on Hippoglossus stenolepis Schmidt, 1904 (Pacific halibut) and Eopsetta jordani (Lockington, 1879) (Petrale sole) (Pleuronectiformes: Pleuronectidae) and parasites geographically isolated in the Atlantic Ocean on H. hippoglossus (Linnaeus, 1758) (Atlantic halibut) and Reinhardtius hippoglossoides (Walbaum, 1792) (Greenland halibut) (Pleuronectiformes: Pleuronectidae) have all been regarded as E. hippoglossi. We confirm that E. hippoglossi sensu stricto is a parasite of the Atlantic halibut, but is accompanied on the same host by a second entobdelline species, E. vanbenedeni n. sp. It differs from the type species, E. hippoglossi in the anatomy of the distal region of the vagina and in the number, distribution and shape of papillae on the ventral surface of the haptor. Parasites from H. stenolepis are morphologically similar to E. hippoglossi on H. hippoglossus (no second species has been found on H. stenolepis), but differences in the number and shapes of haptor papillae support proposal of the parasites from H. stenolepis as a distinct species, E. stenolepis n. sp. Our anatomical studies of these parasites, with the exception of E. pugetensis, have revealed a cluster of several seminal receptacles opening into the ovo-vitelline duct. Previous reports of a single seminal receptacle in E. stenolepis (as E. hippoglossi) and E. steingroeveri are erroneous. Furthermore, reports of the vagina opening proximally into the alleged single seminal receptacle are also incorrect; direct and indirect evidence suggests that in all these parasite species, including E. pugetensis, the vagina opens proximally into the vitelline reservoir. Entobdella squamula from Paralichthys californicus (Ayers, 1859) (California flounder) (Pleuronectiformes: Paralichthyidae) differs from E. stenolepis, E. hippoglossi and E. vanbenedeni in the relative lengths of the accessory sclerites and anterior hamuli. Parasites originally identified as E. squamula from Hippoglossina macrops Steindachner, 1876 (Bigeye flounder) (Pleuronectiformes: Paralichthyidae), caught off the coast of Chile by the Lund University Chile Expedition (1948 – 49), differ from E. squamula in the shapes of the accessory sclerites and are regarded as a new species, E. brinkmanni n. sp. E. aegyptiacus Amer, 1990 must be regarded as a species inquirenda. A key to the 7 valid species of Entobdella recognised by us is included. Entobdella pugetensis differs in the following ways from all other species of Entobdella: the parasite is found on gills (less commonly on skin) of Atheresthes stomias (Jordan & Gilbert, 1880) (Arrowtooth flounder) (Pleuronectiformes: Pleuronectidae); the penis sac has a thick, possibly muscular, wall; the vagina is a relatively wide, short, straight tube with a conspicuous opening; no seminal receptacles have been observed; there are no papillae on the ventral surface of the haptor. We regard these distinctions as sufficient to erect a new genus, Branchobdella n. gen. to accommodate B. pugetensis n. comb. Assimilation of spermatophores in skin-parasitic Entobdella spp. and in the gill parasite B. pugetensis is discussed.


  1. Amer, O.H. (1990) Entobdella aegyptiacus as a new species of monogenean gill trematode of marine fish in Egypt. Veterinary Medical Journal, Giza, 38, 419–427.

    Arthur, J.R. (1984) A survey of the parasites of walleye Pollack (Theragra chalcogramma) from the northeastern Pacific Ocean off Canada and a zoogeographical analysis of the parasite fauna of this fish throughout its range. Canadian Journal of Zoology, 62, 675–684.

    Baster, J. (1765) Opuscula Subseciva, Observationes Miscellaneas de Animalculis et Plantis Quibusdam Marinis, Eorumque Ovariis et Seminibus Continentia. Tom II, Liber II, III. Joannem Bosch, Harlem, pp. 53 – 99 (Liber II), pp. 105 – 150 (Liber III). (In Latin).

    Beverley-Burton, M. (1984) Monogenea and Turbellaria. In: Margolis, L. & Kabata, Z. (Eds), Guide to the Parasites of Fishes of Canada, Part I., Department of Fisheries and Oceans, Ottawa, pp. 7–209.

    Boje, J., Riget, F. & Køie, M. (1997) Helminth parasites as biological tags in population studies of Greenland halibut (Reinhardtius hippoglossoides (Walbaum)), in the north-west Atlantic. ICES Journal of Marine Science, 54, 886–895.

    Brinkmann, A. Jr. (1952a) Reports of the Lund University Chile Expedition 1948 – 1949. 6. Some Chilean monogenetic trematodes. Lunds Universitets Årsskrift, 47, 1–26.

    Brinkmann, A. Jr. (1952b) Fish trematodes from Norwegian waters. I. The history of fish trematode investigations in Norway and the Norwegian species of the order Monogenea. Årbok Universitetet i Bergen. Naturvitenskapelig Rekke, 1, 1–134.

    Bullard, S.A., Payne, R.R. & Braswell, J.S. (2004) New genus with two new species of capsalid monogeneans from dasyatids in the Gulf of California. Journal of Parasitology, 90, 1412–1427.

    Bychowsky, B.E. (1957) Monogenetic Trematodes, their Classification and Phylogeny. Academy of Sciences, USSR, Moscow, Leningrad. (In Russian). (English translation by Hargis W.J. & Oustinov, P.C. 1961. American Institute of Biological Sciences, Washington, 627 pp.).

    Carvalho-Varela, M. & Cunha-Ferreira, V. (1987) Helminth parasites of the common sole, Solea solea, and the Senegalese Sole, Solea senegalensis, on the Portuguese continental coast. Aquaculture, 67, 135–138.

    Cohn, L. (1916) Epibdella steingröveri n. sp. Zeitschrift für wissenschaftliche Zoologie, 115, 460–488.

    Cooper, A.R. (1921) Report of the Canadian Arctic Expedition 1913 – 18. Volume IX: Annelids, Parasitic Worms, Protozoans, etc. Part G-H: Trematoda and Cestoda. Thomas Mulvey, Ottawa, pp. 3–27.

    Crane, J.W. (1972) Systematics and new species of marine Monogenea from California. The Wasmann Journal of Biology, 30, 109–166.

    Egorova, T.P. (2000) Entobdella hippoglossi (Monogenoidea, Capsalidae) from a perch-like fish from the Pacific Ocean and new data on Sessilorbis limopharynx. Parazitologia, 34, 70–74 (In Russian).

    Froese, R.& Pauly, D. (2007) FishBase. Version (02/2007). Available from: (accessed 7 April 2007).

    González, M.T., Acuña, E. & Oliva, M.E. (2001) Metazoan parasite fauna of the bigeye flounder, Hippoglossina macrops, from northern Chile. Influence of host age and sex. Memórias do Instituto Oswaldo Cruz, 96, 1049–1054.

    Gray, P. (1954) The Microtomist’s Formulary and Guide. Constable, London, 794 pp.

    Guberlet, J.E. (1936/1937) Trematodos ectoparasitos de los peces de las costas del Pacifico. Annales del Instituto de Biología. Universidad de México, 7, 457–467.

    Heath, H. (1902) The anatomy of Epidella squamula sp. nov. Proceedings of the Californian Academy of Sciences, 3, 109–136.

    Hendrix, S.S. (1994) Marine Flora and Fauna of the Eastern United States. Platyhelminthes: Monogenea. NOAA Technical Report NMFS 121, U.S. Department of Commerce, Seattle, Washington, 107 pp.

    Hoskins, G.E., Bell, G.R. & Evelyn, T.P.T. (1976) The occurrence, distribution and significance of infectious diseases and of neoplasms observed in fish in the Pacific region up to the end of 1974. Environment Canada, Fisheries and Marine Service, Technical Report No. 609, pp. 1–37.

    International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature, 4th edition, The International Trust for Zoological Nomenclature, 306 pp.

    Kaliszewska, Z.A., Seger, J., Rowntree, V.J., Barco, S.G., Benegas, R., Best, P.B., Brown, M.W., Brownell, R.L. Jr., Carribero, A., Harcourt, R., Knowlton, A.R., Marshalltilas, K., Patenaude, N.J., Rivarola, M., Schaeff, C.M., Sironi, M., Smith, W.A. & Yamada, T.K. (2005) Population histories of right whales (Cetacea: Eubalana) inferred from mitochondrial sequence diversities and divergences of their whale lice (Amphipoda: Cyamus). Molecular Ecology, 14, 3439–3456.

    Kearn, G.C. (1964) The attachment of the monogenean Entobdella soleae to the skin of the common sole. Parasitology, 54, 327–335.

    Kearn, G.C. (1967a) The life-cycles and larval development of some acanthocotylids (Monogenea) from Plymouth rays. Parasitology, 57, 157–167.

    Kearn, G.C. (1967b) Experiments on host-finding and host-specificity in the monogenean Entobdella soleae. Parasitology, 57, 585–605.

    Kearn, G.C. (1970) The production, transfer and assimilation of spermatophores by Entobdella soleae, a monogenean skin parasite of the common sole. Parasitology, 60, 301–311.

    Kearn, G.C. (1971a) The physiology and behaviour of the monogenean skin parasite Entobdella soleae in relation to its host (Solea solea). In: Fallis, A.M. (Ed.). Ecology and Physiology of Parasites, a Symposium. University of Toronto Press, Toronto, pp. 161–187.

    Kearn, G.C. (1971b) The attachment site, invasion route and larval development of Trochopus pini, a monogenean from the gills of Trigla hirundo. Parasitology, 63, 513–525.

    Kearn, G.C. (1985) Observations on egg production in the monogenean Entobdella soleae. International Journal for Parasitology, 15, 187–194.

    Kearn, G.C. (1988) The monogenean skin parasite Entobdella soleae: movement of adults and juveniles from host to host (Solea solea). International Journal for Parasitology, 18, 313–319.

    Kearn, G.C. (1993) Fertilization and the functional anatomy of the germarium in the monogenean Entobdella soleae. International Journal for Parasitology, 23, 901–911.

    Kearn, G.C. (1998) Parasitism and the Platyhelminths, Chapman & Hall, London, 544 pp.

    Kearn, G.C. (2002) Entobdella soleae – pointers to the future. International Journal for Parasitology, 32, 367–372.

    Kearn, G.C. (2004) Leeches, Lice & Lampreys. A Natural History of Skin and Gill Parasites of Fishes. Springer, Dordrecht, The Netherlands, 432 pp.

    Kearn, G.C. (2006) Sub-tegumental ventral fibres surrounding the vaginal opening of Entobdella soleae (Van Beneden et Hesse, 1864) Johnston, 1929 (Platyhelminthes: Monogenea: Capsalidae). Folia Parasitologica, 53, 71–72.

    Kearn, G.C. & Evans-Gowing, R. (1998) Attachment and detachment of the anterior adhesive pads of the monogenean (platyhelminth) parasite Entobdella soleae from the skin of the common sole (Solea solea). International Journal for Parasitology, 28, 1583–1593.

    Kearn, G.C. & Whittington, I.D. (2005) Neoentobdella gen. nov. for species of Entobdella Blainville in Lamarck, 1818 (Monogenea, Capsalidae, Entobdellinae) from stingray hosts, with descriptions of two new species. Acta Parasitologica, 50, 32–48.

    Klassen, G.J., Beverley-Burton, M. & Locke, A. (1989) A revision of Entobdella Blainville (Monogenea: Capsalidae) with particular reference to E. hippoglossi and E. squamula: the use of ratios in taxonomy and key to species. Canadian Journal of Zoology, 67, 1869–1876.

    Lawler, A.R. (1981) Zoogeography and host-specificity of the superfamily Capsaloidea Price, 1936 (Monogenea: Monopisthocotylea). Special Papers in Marine Science No. 6. Virginia Institute of Marine Science and School of Marine Science, College of William and Mary, Gloucester Point, Virginia, USA., 650 pp.

    Llewellyn, J. & Euzet, L. (1964) Spermatophores in the monogenean Entobdella diadema Monticelli from the skin of sting-rays, with a note on the taxonomy of the parasite. Parasitology, 54, 337–344.

    Lyons, K.M. (1973) Scanning and transmission electron microscope studies on the sensory sucker papillae of the fish parasite Entobdella soleae (Monogenea). Zeitschrift für Zellforschung, 137, 471–480.

    Macdonald, S. & Llewellyn, J. (1980) Reproduction in Acanthocotyle greeni n. sp. (Monogenea) from the skin of Raia spp. at Plymouth. Journal of the Marine Biological Association of the United Kingdom, 60, 81–88.

    Mamaev, Yu.L. (1965). Helminths of fish of the Bering Sea. In: Parasitic Worms of Domestic and Wild Animals. Akad. Nauk SSSR, Biol. –Pochven. Inst. 2 Fil. AN SSSR Dal’nevost. Gosudarstvennyi Universitet Vladivostok. (In Russian).

    Margolis, L. (1952) Studies on Parasites and Diseases of Marine and Anadromous Fish from the Canadian Pacific Coast. Unpublished Ph.D thesis, McGill University, Canada, 215 pp.

    Müller, O.F. (1776) Zoologiae Danicae prodromus, seu animalium Daniae et Norvegiae indigernarum characteres, nomina, et synonma imprimis popularium. Havniae. (In Latin). xxxii + 282 pp.

    Oliva, M.E., Castro, R.E. & Burgos, R. (1996) Parasites of the flatfish Paralichthys adspersus (Steindachner, 1867) (Pleuronectiformes) from northern Chile. Memórias do Instituto Oswaldo Cruz, 91, 301–306.

    Price, E.W. (1939) North American monogenetic trematodes. III. The family Capsalidae (Capsaloidea). Journal of the Washington Academy of Sciences, 29, 63–92.

    Robinson, E.S. (1961) Some monogenetic trematodes from marine fishes of the Pacific. Transactions of the American Microscopical Society, 80, 238–261.

    Ronald, K. (1957) The metazoan parasites of the Heterosomata of the Gulf of St. Lawrence. II. Entobdella curvunca sp. nov. (Trematoda: Capsalidae). Canadian Journal of Zoology, 35, 747–750.

    Schram, T.A. & Haug, T. (1988) Ectoparasites on the Atlantic halibut, Hippoglossus hippoglossus (L.), from northern Norway – potential pests in halibut aquaculture. Sarsia, 73, 213–227.

    Sproston, N.G. (1946) A synopsis of the monogenetic trematodes. Transactions of the Zoological Society of London, 25, 185–600.

    Van Beneden, P.–J. (1858) Mémoire sur les Vers Intestinaux. J.–B. Baillière et fils, Paris, 376 pp.

    Van Beneden, P.-J. & Hesse, E. (1864) Recherches sur les bdellodes (hirudinées) et les trématodes marins. Mémoires de l’Académie Royale des Sciences, des Lettres et des Beaux-arts de Belgique 34, 1–142.

    Whittington, I.D., Deveney, M.R., Morgan, J.A.T., Chisholm, L.A. & Adlard, R.D. (2004) A preliminary phylogenetic analysis of the Capsalidae (Platyhelminthes: Monogenea: Monopisthocotylea) inferred from large subunit rDNA sequences. Parasitology, 128, 511–519.

    Whittington, I.D. & Kearn, G.C. (1991) The adhesive attitudes of some gill-parasitic capsalid monogeneans. Journal of Helminthology, 65, 280–285.

    Yamaguti, S. (1963) Systema Helminthum. Vol. IV Monogenea and Aspidocotylea. Interscience Publishers, New York, London, 699 pp.